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Funding: The synthetic work for this paper was 151 dams with capacities larger than 2 MegaWatts (MW) in the western Amazon during the
supported by the Science for Nature and People next two decades [2]. Most of these dams would be built in the Andes Mountain where steep
(SNAP) sponsored by the National Center for
topography facilitates the creation of deep storage reservoirs with high hydraulic head. Six
Ecological Analysis and Synthesis (NCEAS), the
Wildlife Conservation Society (WCS) and the dams, planned for construction on major Andean tributaries with high suspended sediment
Nature Conservancy (TNC). At WCS we thank concentrations, are of particular concern because they will be the largest and farthest down-
Cristián Samper, John Robinson, Julie Kunen, stream storage dams on their respective tributaries [2]. Together, these impoundments could
Mariana Varese, Mariana Montoya, Carlos Durigan, have major impacts on the hydrology, geomorphology, biogeochemistry, biodiversity and pro-
Guillermo Estupiñán, Micaela Varese, Natalia Piland
ductivity of the Amazon River system, affecting human livelihoods and well-being from the
and Sofia Baca; for SNAP workshop support we
thank Charo Lanao; at TNC Craig Groves and Peter
headwaters to the estuary. Significant changes are expected both downstream and upstream of
Kareiva; at NCEAS Frank Davis and Lee Ann these dams.
French. We thank D. Kasper, J.R.P. Peleja, E.M. Based on their mineralogy, Gibbs [3] concluded that most sediments carried by the Ama-
Nakazono, A.P. Souza, J.B. Rocha and D.R. zon River are derived from the Andes (Fig 1). Current estimates suggest that 93% of all sedi-
Dietrich for help during the collection and analysis ments in the Amazon River system are derived from this source [4]. High correlations
of fish and hair for mercury at Balbina Reservoir
between total suspended sediments (TSS) and particulate phosphorus and nitrogen concentra-
and orientation. We also thank the Museu Paraense
Emilio Goeldi (MPEG) and the Instituto Nacional de tions in these rivers [5] indicate that most sediment-bound nutrients in the river system are
Pesquisas da Amazônia (INPA) for their support. also derived from the Andes. The damming of major tributaries draining the Andes could,
For fisheries data we are indebted to the Dirección therefore, reduce the supply of both sediments and nutrients to the Amazon lowlands (Fig 1).
Regional de la Producción (DIREPRO), Peru. SNAP While dam building is only beginning in the Amazon, dams have already caused significant
funding was provided by the David and Lucile
declines in the sediment loads of other large river systems with major consequences for fluvial
Packard Foundation (Grant # 2013-38757 &
#2014-39828), Ward Woods (Grant # 309519), the
ecosystems and human populations downstream [6–10]. Large reductions in sediment loads
Wildlife Conservation Society (WCS) and The following impoundment have resulted in rapid downstream channel erosion with decreases in
Nature Conservancy (TNC). Field data reported for bed elevations, changes in channel width and the loss of riparian habitats and vegetation [7,
the first time was supported by the Gordon and 10, 11–14]. In rivers where sediment supplies are not replenished by downstream sources,
Betty Moore Foundation (Grant 500) and the John these impacts have extended to lowland floodplains and deltas where the loss of residential
D. and Catherine T. MacArthur Foundation (Grant
areas and farmlands due to increased flooding and subsidence is now a global concern [9].
84377). The Brazilian National Research Council
(CNPq) provided a productivity grant to B.R. Reductions in the supply of riverine nutrients to downstream floodplains, deltas and coastal
Forsberg (Grant 309636/2011-6). J.M. Melack areas have led to losses in soil fertility, accompanied with increased use of artificial fertilizers
received support from the US Department of [8] and declines in aquatic primary production and fish yield [7, 8, 15, 16]. The changes
Energy (Contract No. DE-0010620), NASA and a expected below the new Andean dams will depend on the amount of sediment and nutrients
Fulbright fellowship. From the above institutions,
retained by these impoundments and the hydrological and geomorphological dynamics of the
past or present, we thank Avecita Chicchón, Adrian
Forsyth, Rosa Lemos da Sá and Enrique Ortiz.
river systems below. Downstream inputs from tributaries and channel exchange processes in
the foreland and lowland regions [4, 17–20] may replenish, in part, the supply of sediments
Competing interests: The authors have declared
and nutrients retained by the dams before the rivers reach the lowland floodplains (Fig 1).
that no competing interests exist.
However, if the initial decline in sediment and nutrient supplies below the dams are large and
a significant part of this reduction propagates downstream, lowland environments are likely to
be affected.
The seasonal flood-pulse of the Amazon River plays a fundamental role in maintaining the
diversity and productivity of lowland floodplain environments [21, 22]. Inputs of river-borne
nutrients during seasonal floods maintain the fertility of lowland floodplain soils [23–25] and
the productivity of alluvial wetlands [26–28]. The dynamic interplay between floodplain
topography and river flood cycle [29, 30] creates a complex mosaic of floodplain environments
with varying flood dynamics and a diversity of aquatic flora and fauna adapted to these condi-
tions [1, 31–36]. The production dynamics and phenology of this biota are synchronized to
local inundation patterns [21, 37, 38]. Fish production and yield are linked to flood dynamics,
with the highest yields occurring 1–2 years after the largest floods [40–41].
Variations in water discharge through hydroelectric turbines are normally limited by dam
managers in order to stabilize power generation [11, 42]. This operational norm can have a
large impact on the flood-pulse below the dam, reducing maximum stage heights and flooded
areas, increasing minimum stage heights and flooded areas and altering the spatial pattern of
Fig 1. Amazon basin, showing main hydrological and geomorphological features and locations of proposed Andean dams. Locations of river
gauging stations and lowland dams mentioned in the text indicated. Topography derived from the Shuttle Radar Topographic Mission Digital Elevation
Model, National Aeronautics and Space Administrations, USA (SRTM-DEM, NASA).
https://doi.org/10.1371/journal.pone.0182254.g001
inundation across the floodplain [11, 43, 44]. If similar changes occur below the Andean dams,
they could have profound effects on the flora, fauna and human populations occupying these
regions, including a reduction in floodplain fertility and fish yield, the permanent flooding
and mortality of low lying vegetation and the large-scale disruption of plant phenology and
growth cycles [1, 36, 45].
A different suite of impacts is expected upstream of the Andean dams, as fluvial ecosystems
are transformed into reservoirs. The transformation of a flowing river and its accompanying
upland basin into a large lake results in the destruction of terrestrial vegetation and major
changes in the structure and functioning of the aquatic ecosystem [1, 46]. The generally well
mixed riverine water column becomes thermally stratified in the reservoir. Fine sediments and
sands, which were suspended in the river, decant in the reservoir, causing the siltation of ben-
thic environments [13]. As inundated terrestrial vegetation dies and decomposes, dissolved
oxygen levels fall and significant amounts of dissolved organic matter, nutrients and green-
house gases (GHGs:CO2 and CH4) are released to the overlying water and atmosphere [47–
49]. These conditions favor the methylation of dissolved inorganic mercury (MeHg), originally
present in the inflowing river, and its bioaccumulation in fish and other aquatic organisms
[50, 51]. GHGs and MeHg produced in reservoir bottom waters are also exported to the fluvial
ecosystem below the dam, contributing to increased GHG emissions and mercury contamina-
tion in these regions [47, 48, 52, 53]. Increased nutrient concentrations from internal and
external loading together with improved light penetration generally result in elevated levels of
primary production and fish yield in reservoirs, which can lead to the development of impor-
tant local fishing industries, especially in tropical reservoirs [54, 55]. Elevated fish production
is expected in the new Andean reservoirs due to large inputs of nutrients from Andean
tributaries [56]. This is likely to bring economic benefits to these regions but may also promote
mercury contamination in the populations that consume these fish.
Several recent attempts have been made to evaluate the potential impacts of hydropower
development on the Amazon [1, 2, 9, 36, 57, 58] and other large tropical river ecosystems [9, 57,
59]. These analyses have focused on aquatic and terrestrial biodiversity [36, 57, 59], sediment
dynamics [9] or integrated multiple impacts [1, 2, 58]. They have generally been qualitative
analyses, at most comparing regional distributions of dams with maps of species richness or
habitats [57, 59]. The exception was a recent analysis of Labtrubesse et al. [58], which used spa-
tial indexes, based on quantitative estimates of current land use cover, sediment yield, hydrolog-
ical variability, geomorphological variability, and fluvial connectivity, to evaluate current and
future vulnerability to hydropower development across the Amazon Basin. While this analysis
attributed the highest vulnerability to basins draining the Andean headwaters, the exact nature
and magnitude of these impacts were not examined. Quantitative predictions of these impacts,
based on mechanistic and process-based associations with dam construction, are needed.
Here we use historical data from Amazonian rivers and reservoirs, together with mechanis-
tic scenarios to examine the potential impacts of six planned Andean dams on Amazon fluvial
ecosystems. We consider impacts both above and below the dams, including 1) the reduction
in downstream sediment supply, 2) the reduction in downstream nutrient supply, 3) changes
in downstream flood pulse, 4) changes in upstream and downstream fish yield, 5) reservoir sil-
tation, 6) greenhouse gas emissions above and below the dams, and 7) mercury contamination
above and below the dams. The analysis provides quantitative predictions of the magnitude
and extent of these impacts along with descriptions of the methods and relationships used to
generate them.
Fig 2. Locations of planned Andean dams and the Loreto fishing territory. Impacted tributaries, relevant gauging stations and key
geomorphological features are indicated. Topography derived from NASA, SRTM-DEM [60].
https://doi.org/10.1371/journal.pone.0182254.g002
0:19ðVIÞ
T ¼ 100 ½0:97 ð1Þ
https://doi.org/10.1371/journal.pone.0182254.t001
the ratio of these values. The expected decline in sediment supply for the entire Amazon due
to the dams was estimated from the % reduction in Andean yield, assuming the Andes
accounts for 93% of the basin-wide yield [4].
Amazonian yield (approximately 0.93 [4]). Total dissolved nutrient yields for the Amazon
were estimated from the product of the average discharge of the Amazon River at its mouth
(205,000 m3 s-1 [69]) and the average concentration of total dissolved nutrients determined by
the CAMREX Project for Amazon tributaries (TDN = 325 mg m-3; TDP = 39 mg m-3).
The current supply of PP to the central floodplain and delta plain regions was estimated by
multiplying the combined annual accumulation of sediment in these regions [19] by the aver-
age phosphorus content of Amazon River sediments (0.08%) [5]. The input of PN was esti-
mated in a similar way using the average N content (by weight) of river sediments along the
central Amazon floodplain determined by the CAMREX project (0.11%, B. Forsberg, unpub-
lished data). The reduction in these supplies following dam construction was presumed equiv-
alent to the basin-wide drop expected for sediments.
Fig 3. Locations of A) Balbina Dam on the Uatumã River and B) Tucurui Dam on the Tocantins River, showing reservoirs and locations of
downstream gauging stations. Cachoeira Morena and Tucurui gauging stations are indicated below the dams. Station locations from the Brazilian
National Water Agency, ANA [70]. Reservoir area derived from NASA, SRTM-DEM and LANDSAT imagery [60].
https://doi.org/10.1371/journal.pone.0182254.g003
for the territory exploited by commercial and riparian community fishermen in the Loreto
region of Peru (Fig 2). This area is composed of a complex matrix of river channels, and sea-
sonally inundated alluvial floodplains. Fish production in the region is sustained by high levels
of primary production on floodplains supported by river-borne nutrients. Data for annual fish
yield were obtained from DIREPRO (Dirección Regional de Producción, Peru) for the main
cities along the Amazon, Ucayali and Marañón floodplains between 2004 and 2011 and was
organized by fishing zone, fishing fleet and species (S1 Table). Total annual yield included all
fish caught in the fishing zones located in the fishing territory delimited in Fig 2.
Daily flooded areas for the Loreto fishing territory were estimated for a 12 year period
(1998–2009), using the MGB-IPH large-scale numerical hydrological model, that was devel-
oped and validated specifically for the Amazon Basin [71, 72]. Maximum annual flooded areas
(MxFA) and minimum annual flooded areas (MiFA) for the period 1999–2011, that included
5 years before the beginning of the fish yield record and 2 years after the simulated data set,
were estimated from a linear regression between simulated daily flooding in the Loreto fishing
territory and daily stage height measured at Iquitos, (SENAMI, Peru) (S1 Table).
Inter-annual variation in fishing effort has been shown to be an important predictor of fish
yield in the Amazon [73]. While precise information on fishing effort was unavailable for the
major fishing zones [74, 75], the total number of fishing zones (FZ) varied between years and
was used as a proxy for fishing effort in the analysis. Historic variations in fish yield, fishing
zones, minimum flooded area and peak flooded area for the Loreto Region are provided in S1
Table. The influence of MxFA, MiFA and FZ on annual fish yield was investigated with simple
and multiple linear regression analyses, using time lags of 0–5 years to evaluate the delayed
effects of flooding, as suggested by Welcomme [39]. The 10 year average maximum inundated
area was then used together with the best regression model to predict the effect of proportional
reductions in mean maximum flood extent, due to impoundment, on fish yield.
Reservoir sedimentation
The information necessary to estimate reservoir sedimentation rates was available for the
Angosta del Bala, Pongo de Manseriche, Rositas and Inambari reservoirs. Storage volumes
were estimated from specified dam locations and reservoir surface heights [78], using the
SRTM—DEM [60] (Table 1). All reservoirs will be a deep and steep sided, with average depths
ranging from 19–103 m. The number of years required to completely fill each reservoir was
estimated by dividing the storage volume by the volume of sediments delivered to the system
annually. The volume of suspended sediments reaching the reservoir was estimated by divid-
ing the annual sediment discharge (by weight) near the proposed dam site [18, 61] (B. Fors-
berg, unpublished) by the bulk density of recently deposited Amazon sediments, estimated by
Devol et al.[79] to be 1,396 kg m-3. The additional contribution of bedload to sediment volume
was estimated assuming that bedload transport was 10% of total sediment discharge [62] and
that the bulk density of bed material was similar to that of cobblestones, estimated by Carling
and Reader [80] to be 1890 kg m-3.
were estimated using the following empirical relationships, derived from data for 85 globally
distributed reservoirs [82]:
LogðCO2 flux þ 400Þ ¼ 3:06 0:16LogðageÞ 0:01Lat þ 0:41LogðDOCÞ ð3Þ
where,
DOC = dissolved organic carbon concentration in the reservoir, mgC L-1,
DOCLoad = DOC loading to the reservoir, mgC m-2 d-1,
age = years since impoundment,
mean depth = the mean depth of the reservoir in meters, and
Lat = mean latitude of the reservoir
External DOC loading was estimated from the product of tributary inflow and DOC con-
centration divided by reservoir area. DOC concentrations were obtained from the ORE-HY-
BAM project (www.ore-hybam.org) or from an empirical relationship between DOC and
elevation developed for Andean tributaries [83]. External DOC loading and background DOC
concentrations were assumed to be constant over time. Additional internal DOC loading,
derived from the decay of inundated terrestrial vegetation, was assumed to decline gradually
following impoundment as the terrestrial carbon stock diminishes. The initial areal carbon
stock of terrestrial vegetation in all reservoir areas was assumed equivalent to that estimated by
Vega et al. [84] for the Inambari Reservoir, 30,300 tons C km-2. Total terrestrial carbon stock
was estimated by multiplying this value by the projected surface area of each reservoir. This
carbon stock and internal DOC loading was assumed to decline over time at an exponential
decay rate of 0.23 y-1, similar to the rate of decline in emissions observed at Petit Saut Reser-
voir, French Guyana, after impoundment [47]. This variable loading rate was used, together
with the estimates of upstream loading and background DOC, to calculate the change in DOC,
DOCLoad and greenhouse gas emissions over time. Methane emissions were converted to
CO2-equivalents (CO2e) assuming a 100 year global warming potential (GWP) for methane of
34 [85].
Significant quantities of greenhouse gases are also released downstream from dams [47, 48,
86, 87]. These fluxes include both degassing at the turbine outflow and diffusive emissions
from the downstream river. The total downstream flux has been shown to be proportional to
the power generating capacity of a dam [87] and to its turbine discharge [88]. The downstream
emissions of CO2-equivalent carbon for Balbina and Petit Saut dams, normalized to turbine
discharge, were 22.6 and 26.9 gC-CO2e m-3, respectively, with an average value of 24.8 gC-
CO2e m-3 [88]. This average value was multiplied by the annual discharge of each reservoir to
estimate the initial CO2 equivalent carbon emission expected downstream of the dams. Since
downstream emissions are fueled, in large part, by internal carbon loading to a reservoir [47],
an exponential decay rate of 0.23 y-1 was also applied to these initial values to simulate the
expected decline over time.
In order to compare emission characteristics between dams and between these planned
hydroelectric facilities and alternative energy sources, we estimated Carbon Emission Factors
(CEFs) for each dam. These factors were estimated for each dam by dividing the total annual
CO2 equivalent carbon emission in tons C-CO2e, including both reservoir and downstream
fluxes, averaged over the first 30 years of dam operation, by the total annual power generation
of the dam in MWhrs.
Mercury dynamics
Empirical models have been developed for tributaries of the Amazon relating mercury levels
in fish and human hair to trophic level, pH, DOC and wetland densities [89–91]. However,
these relationships vary geographically [92], and there are insufficient data for Andean
tributaries to develop specific relationships for this region. The dynamics of mercury in reser-
voirs is also different than that found in free flowing river systems, due to the anoxic condi-
tions, expected following impoundment, that promote high rates of mercury methylation and
subsequent biomagnification. Relationships between the levels of MeHg in water or fish and
terrestrial carbon stocks or percent flooding relative to volume and have been developed for
some north temperate reservoirs [93, 94], but similar relationships are unavailable for tropical
systems. Due to the lack of mechanistic models for predicting Hg dynamics in the Andean
dams, we were limited to examining the temporal variation of Hg contamination in Balbina
(Fig 3), the only Amazonian reservoir with a consistent historical time series of mercury mea-
surements. There was no gold mining or other anthropogenic sources of mercury, besides
regional atmospheric inputs, in the drainage basin upstream of Balbina Dam, during the
period considered. Mercury present in the system is presumed to be predominantly of natural
origin. Historical data for Balbina Reservoir included measurements of total mercury in Cichla
spp., the main fish predator in the system (B. Forsberg, unpublished data), and total mercury
in the hair of fish-eating human populations that historically exploited this resource [95]. Fish
of varying sizes were collected between 1992 and 2003, while Hg levels in hair were obtained
for the period of 1995–2000 [95]. For the unpublished fish data, boneless and skinless samples
of dorsal muscle were collected and stored frozen until analysis. Following digestion these
samples were analyzed for total mercury by cold vapor atomic absorption spectrophotometry
[96, 97]. Mean Hg concentrations in fish were normalized to a standard size.
Results
Impact on the downstream sediment supply
The combined reduction in sediment discharge expected for all six rivers was estimated at 894
x 106 tons y1 (Table 2). The estimated area of all Andean highlands with elevation >500 m and
the total area of these highlands drained by the six dam basins were 628,000 km2 and 436,000
km2, respectively. The percent of the Andean highlands drained by the six dam basins and the
Table 2. Predicted reduction in sediment discharge below six planned Andean dams following
impoundment.
Dam Predicted reduction in
sediment discharge,106 t y-1
Angosto del Bala 243a
TAM 40 272b
Pongo de Aguirre 57b
Pongo de Manseriche 166b
Rosita 138a
Inambari 18c
TOTAL 894
Estimates assume that 100% of suspended and bedload sediments are retained by all dams.
a
[18].
b
[61].
c
B. Forsberg, unpublished data.
https://doi.org/10.1371/journal.pone.0182254.t002
expected decline in Andean sediment yield, represented by the above value, is therefore 69%.
Since export from the Andes accounts for approximately 93% of all sediment yield in the Ama-
zon basin [4], the construction of these six dams is expected to result in a 64% reduction of in
the basin-wide sediment supply.
Table 3. Predicted reduction in nutrient fluxes below six Andean dams after impoundment.
Dam Current fluxes With dam Reduction % change
106 tons y-1 106 tons y-1 106 tons y-1
TP TN TP TN TP TN TP TN
Angosto del Bala 0.177 0.233 0.002 0.021 0.175 0.212 -99 -91
TAM 40 0.201 0.280 0.005 0.043 0.196 0.237 -98 -85
Pongo de Aguirre 0.044 0.080 0.003 0.030 0.041 0.049 -92 -62
Pongo de Manseriche 0.125 0.195 0.006 0.051 0.119 0.144 -95 -74
Rosita 0.100 0.124 0.000 0.003 0.100 0.121 -100 -98
Inambari 0.015 0.031 0.002 0.016 0.013 0.016 -88 -50
TOTAL 0.662 0.944 0.019 0.164 0.643 0.780 -97 -83
Estimates assume 100% retention of particulate fractions and no retention of dissolved fractions. Sources for data used in calculations: [18, 61, 5], B.
Forsberg, unpublished data, CAMREX PROJECT (http://dx.doi.org/10.3334/ORNLDAAC/904).
https://doi.org/10.1371/journal.pone.0182254.t003
Fig 4. Changes in average monthly stage heights and inundation periods for stage heights of the Uatumã River below Balbina dam (A
and B, respectively) and of the Tocantins River below Tucurui dam (C and D, respectively), following impoundment. Pre-construction
periods (blue line) = 1973–1982 for Balbina and 1969–1975 for Tucurui; post impoundment periods (red line) = 1991–2011 for Balbina and 1985–
2014 for Tucurui. Stage data obtained from Brazilian National Water Agency, ANA [70]. Standard error bars indicated.
https://doi.org/10.1371/journal.pone.0182254.g004
flooding patterns in these areas. Low lying areas, below a stage height of 325 cm on the
Uatumã floodplain and below 200 cm on the Tocantins floodplain, that were seasonally dry
before impoundment are now almost permanently flooded, while upland portions of the
Uatumã floodplain, above a stage height of 475 cm, that were seasonally flooded at high water
are now almost permanently dry. If we assume a linear relationship between river stage height
and inundated area for the Uatumã floodplain, the observed decline in mean stage height rela-
tive to PSR at peak high water, would have represented a 37% reduction in peak flooded area.
Changes in the general pattern of inundation on the Uatumã (Fig 4B) and Tocantins flood-
plains (Fig 4D) also differed. Areas above a stage height of ~430 cm on the Uatumã floodplain
all had shorter flood periods while areas below this level had longer flood periods. Flood peri-
ods at all elevations except 430 cm changed significantly. Flood periods also changed at stages
below ~670 cm on the Tocantins floodplain, with shorter flood periods occurring above ~280
cm and longer periods below this elevation. No significant changes in flood period were
encountered at stage heights above 670 cm.
Reservoir sedimentation
Predicted sediment filling times for the four Andean dams ranged from 106 years for Angosta
del Bala to 6240 years for Manseriche Reservoir (Table 6). The implications of these results for
the reservoir environment are considered below.
Table 4. Regression models: Influence of inundation and fishing effort on annual fish yield in Loreto, considering the effect of different time lags
after inundation.
Regression model Time lag (years after inund.)/r2 Equation parameters, 2 year lag
0 1 2 3 4 5 a sa b sb c sc
TY = a+b*MxFA - 0.07 0.75* 0.26 0.07 - -25,723 9,655 1.50 0.32
TY = a+b*MiFA 0.15 - - 0.03 - -
TY = a+b*MxFA+c*FZ - - 0.70* 0.14 - - ns ns 1.50 0.35 ns ns
TY = a+b*MiFA+c*FZ 0.01 - - - - -
YFFZ = a+b*MxFA - 0.06 0.83* 0.28 0.14 - -22,755 6,288 1.20 0.21
YFFZ = a+b*MiFA 0.19 - - 0.02 - -
TY = total yield, tons y-1; YFZ = yield for regularly exploited fishing zones, tons y-1; FZ (fishing effort) = number of fishing zones; MxFA = maximum flooding
area, km2 and MiFA = minimum flooding area, km2; Regression parameters (a, b and c) only shown for significant regressions with 2 year lag; r2 = %
variance in yield explained by regression; s = standard error of parameter; ns = not significant.
* indicates significance at p< 0.05 level.
https://doi.org/10.1371/journal.pone.0182254.t004
Fig 5. Relationship between total annual fish yield and the maximum flooded area two years earlier for the Loreto Region of Peru.
Flooded areas simulated using the MGB-IPH large scale numerical hydrological model, developed specifically for the Amazon Basin [71, 72].
Annual fish yields obtained from DIREPRO (Dirección Regional de Producción, Peru).
https://doi.org/10.1371/journal.pone.0182254.g005
Manseriche Dam. Average emissions from the reservoirs were lowest for Rositas and highest
for Manseriche Reservoir. Mean emissions downstream of the reservoirs were also lowest for
Rositas and highest for Manseriche, and averaged 17% of mean total emissions for all dams.
Carbon Emission Factors based on mean total emissions were lowest for Inambari and highest
for Manseriche Dam with an arithmetic average of 0.091 tons C-CO2e MWhr-1 and a MW
weighted average of 0.139 tons C-CO2e MWhr-1 for all dams.
Predicted annual emissions above and below all dams declined during the first 10 years of
operation and then remained relatively stable, as illustrated in results from Manseriche Dam
(Fig 7). Emissions from Manseriche Reservoir were predicted to decline from an initial high of
33.2 X 106 to 2.5 X 106 tons C-CO2e y-1 30 years after impoundment. Downstream emissions
were significantly lower, declining from an initial high of 3.9 X106 to 0.4 X106 tons C-CO2e y-1
over the next 30 years. Total emissions declined from an initial high of 37.1 X 106 tons C-CO2e
y-1 to a rate of 2.8 X 106 tons C-CO2e y-1 after 30 years. The total cumulative CO2-equivalent
carbon emission for Manseriche Dam estimated for the first 30 years after impoundment
was 237 X 106 tons C-CO2e, with 88% released from the reservoir surface and 12% released
Fig 6. Predicted decline in annual fish yield for the Loreto Region due to a reduction in maximum annual flooded area following
impoundment. Relation simulated using percentage changes in average maximum flooded area (MxFA) and regression model: Y = 1.5(MxFA)
− 25,723, with a 2 year lag.
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downstream of the dam. The mean daily areal CO2-equivalent carbon emission rates estimated
for Angosta del Bala, Manseriche, Rositas and Inambari reservoirs for the 30 period were
2,660, 3,090, 2,020 and 3,790 mg C-CO2e m -2 d -1, respectively, with an average value for all
reservoirs of 2,890 mg C-CO2e m -2 d -1.
https://doi.org/10.1371/journal.pone.0182254.t005
https://doi.org/10.1371/journal.pone.0182254.t006
Mercury dynamics
Average size normalized mercury levels in Cichla spp. from Balbina Reservoir increased from
a low of 0.15 μg g-1 in 1992 to a peak of 0.65 ppm in 1997, 10–11 years after impoundment,
and then declined to 0.32 μg g-1 by 2003 (Fig 8). Average mercury levels in the hair of local
human populations who consumed these fish as their main protein source followed a similar
trend (Fig 8) [95], rising from an initial value of about 4.3 μg g-1 in 1995 to a peak of 7.5 μg g-1
in 1999 and then declining to 5.6 μg g-1 by 2000.
Discussion
Impact on the downstream sediment supply
Theoretical and historical evidence from other impounded rivers indicates that the 900 x 106
tons y-1 decrease in sediment discharge expected if the six new Andean dams are built will
have serious consequences for fluvial and riparian ecosystems downstream from these sites
[98]. The nature and timing of these impacts and the extent to which they propagate through
the fluvial ecosystems will depend on the hydrologic and geomorphic characteristics of the
river system, which vary from the Andean forelands to the Amazon Delta (Fig 1). Based on
results from the Amazon [20] and other river systems [11,14], the largest impacts are likely to
occur in the foreland reaches closest to the dams, where large reductions in sediment supply
and moderate reductions in peak water discharge are expected to result in erosion and incision
of channel beds and decreased rates of channel migration. The gradual coarsening or stripping
of finer sediments is expected to produce armored, rocky beds in the cobblestone/sand reaches
closest to the dams and coarse sand beds throughout most of the foreland basin reaches.
Williams and Wolman [11] evaluated increases in channel-bed depth (incision) down-
stream from 6 storage reservoirs along the Colorado and Missouri rivers, comparable in size to
Table 7. Estimated average CO2-equivalent carbon emissions for the reservoirs and downstream reaches of 4 planned Andean dams during the
first 30 years of operation.
Dam MW 30 year mean emission, 105 tons C-CO2e y-1 Emission Factor
Reservoira Downstreamb Total tons C-CO2e MWhr-1c
Angosta del Bala 1600 6.4 3.8 10.2 0.072
Pongo de Manseriche 4500 71.8 9.3 81.1 0.206
Rositas 400 1.2 0.6 1.8 0.051
Inambari 1500 1.5 2.9 4.4 0.034
Total 8,000 80.9 16.6 97.5 mean 0.091
a
Reservoir emissions estimated following [82].
b
Downstream emissions estimated following [88].
c
Carbon Emission Factors estimated from mean total CO2 equivalent emission and specified power generation.
https://doi.org/10.1371/journal.pone.0182254.t007
Fig 7. Predicted emissions of CO2-equivalent carbon upstream and downstream of Manseriche Dam during the first 30 years of
operation. Reservoir emissions estimated following [82]. Downstream emissions estimated following [88].
https://doi.org/10.1371/journal.pone.0182254.g007
those planned for the Grande and Inambari Rivers. The authors analyzed post-impoundment
variations in channel morphometry and developed empirical models to predict the maximum
increase in channel depth at each cross-section, dmax (m), as well as the time it would take to
reach 50% and 95% of this maximum value, T0.5 and T0.95, respectively. Average dmax values
below these dams ranged from 1.4–4.9 m with T0.5 values ranging from 4–13 years. The
remaining change was slower due to an increase in particle sizes and the development of
armored beds with average T0.95 values from 83–246 years. The length of these degraded
reaches and the rate at which they expanded downstream varied between sites, depending on
inputs of sediments from tributaries downstream from the dams [14]. After twenty years they
extended from 15 to greater than 120 km.
The extent of channel incision below larger dams, like those planned for the Ucayali, Mar-
añón and Huallaga rivers, is expected to differ is some respects. Bed elevations in the Nile
River below the Aswan High Dam were found to decrease 0.3–0.7 m after impoundment [99]
but it was not known how far downstream these changes extended. Dai and Lui [10] found
much larger changes in the channel of the Yangtze River downstream from the Three Gorges
Dam with evidence of incision depths ranging from 1–2.5 meters and extending 670 km below
Fig 8. Historical variation of mercury levels in Cichla spp.(blue line) and in the hair of fish-eating residents (red line) from Balbina
Reservoir following impoundment. Data from [95] and B. Forsberg, unpublished.
https://doi.org/10.1371/journal.pone.0182254.g008
the impoundment. The degree of incision below larger dams thus appears to be similar to that
below the smaller dams but can extend much farther downstream.
As bed elevations in foreland rivers decline, river surface levels are expected to fall, reducing
inputs of water, sediments and nutrients to adjacent floodplain environments during seasonal
floods. In addition to reducing floodplain fertility, this is expected to alter spatial patterns of
inundation, critical to floodplain plants [21, 35]. Reduced connectivity between channels and
floodplains could also affect the mobility of human populations and the movements of fish
and other aquatic organisms that migrate between these environments to feed or reproduce.
Analyzing the temporal dynamics of river channels and floodplain geomorphology across
the Amazon basin, Constantine et al. [20] demonstrated that the rate of channel change, the
rate of channel avulsion and the frequency of oxbow lakes were all positively correlated to
width normalized sediment discharge and that the largest effects occurred in the Amazon fore-
land regions. Based on their findings, we expect the massive reductions in sediment discharge
predicted for the foreland reaches below the Andean dams to result in a major reduction in the
rates of channel migration and avulsion and in the frequency of oxbow lakes along these chan-
nels. Based on studies of forest dynamics in this same region [100], we expect the decline in
channel migration to result in a general decrease in plant diversity in both terrestrial and
aquatic environments. Increased stability and reduced inundation of the floodplains along
these reaches is also likely to promote agricultural development with accompanying increases
in deforestation. Rates of erosion and deposition in both channels and alluvial wetlands are
also expected to decline [20], reducing the fertility and sediment storage capacity of these
environments.
The Sub-Andean forelands have trapped about 50% of all the sediment leaving the Andes
throughout the last ~10 M years and continue to do so to varying degrees in the era of instru-
mental sediment measurements [63, 61]. In the Beni River, for example, almost half of the 243
x 106 tons y-1 of sediment passing by the Angosto del Bala gorge is currently deposited in the
subsiding foreland basin [18, 98]. Similar levels of storage have been reported in the forelands
of the Grande and Marañón Rivers [18, 63]. A decrease in the rates of sedimentation in these
foreland basins could therefore have a significant effect on the sediment balance of these
regions.
The extent to which reduced sediment loads and their effects propagate through the low-
land river system will depend on the magnitude of sediment inputs from downstream tributar-
ies and lowland channel-floodplain exchange processes. Results from other large river systems
provide some insight into the expected trends. In the Yangtze River below the Three Gorges
Dam, which receives sediment inputs from minor tributaries and channel erosion, sediment
discharge was 40% lower than pre-impoundment levels 1200 km below the dam [10]. In the
Nile River below the Aswan High Dam which has no significant downstream tributaries, sedi-
ment loads were still 80% below pre-impoundment levels 965 km downstream from the
impoundment [6]. In the Mississippi River, which has dams retaining sediments along all of
its major tributaries, the median sediment load just above its delta is currently 73% below
pre-impoundment levels [101], despite large tributary inputs. A different pattern might be
expected below the Andean dams where most tributaries are still largely unregulated. The fore-
land and lowland tributaries below the six Andean dam sites drain about 31% of the Andean
highlands and presumably account for about 29% of the current sediment load of the Amazon
main channel, assuming the Andes yield is 93% of the total [4]. Inputs from rivers draining the
Guyana and Brazilian Shields could increase this proportion by up to 7% [4]. Downstream sed-
iment sources can therefore expected to maintain the sediment load of the lower Amazon
main channel at most at 36% of its pre-impoundment level.
Channel erosion and deposition have been shown to play a major role in the sediment
dynamics of the lowland Amazon. A detailed sediment balance for the 1560 km reach of the
Amazon mainstem between São Paulo de Olivença and Obidos [19] (Fig 1) demonstrated that
sediment exchanges due to bank erosion (1570 x106 tons y-1), bar deposition (380 x106 tons
y-1) and floodplain sedimentation (1690 x106 tons y-1) were of same order of magnitude as the
total tributary input (1448 x 106 tons y-1). The cumulative result of tributary inputs and chan-
nel exchange processes along the entire reach was a net reduction of at least 200 x106 tons y-1
in the downstream sediment flux, which was deposited along the central Amazon floodplain.
An additional 300–400 x 106 tons y-1 are currently deposited on the Delta plain below Obidos
[19]. Based on the relationships reported by Constantine et al. [20], we expect the magnitude
of channel exchange processes to decline as main channel sediment loads decrease following
impoundment while the lowland floodplain and delta regions should continue to act as a net
sink for sediments in the region.
If the initial drop in sediment discharge caused by the Andean dams is replenished only by
downstream tributaries and the forelands and the mainstem lowlands continue acting as net
sediment sinks, the sediment load of the Amazon mainstem is expected to fall by at least 64%
and this could have major consequences for the central floodplain and delta regions. Channel
bed scouring is expected to reduce hydrological connectivity between channels and floodplains
and alter the inundation regimes in floodplain lakes and wetlands, with associated impacts on
aquatic flora and fauna [21, 35, 37]. The reduction of sediment concentrations will lead to a
large reduction in the quantities of suspended sediment decanted into the floodplain via flood-
plain channels and diffuse overbank flow [19]. The mainstem lowlands are the most densely
populated regions in the Amazon and the most important for agriculture and fish production.
Decreased inputs of these nutrient rich sediments could alter the morphology and hydrology
of these depositional environments [9, 20] and potentially reduce their agricultural and aquatic
productivity.
the reduction in nutrient levels. Historic declines in fish yield in the Nile River and coastal areas
downstream from the Aswan High Dam have also been attributed to lower levels of nutrients
and aquatic primary production associated with nutrient retention by the reservoir [7, 8].
The reduction in the N:P ratio of the downstream nutrient supply, due to the selective
retention of P-rich sediments following impoundment, could also affect the patterns of nutri-
ent limitation in lowland floodplain plants, intensifying the tendency toward N-limitation
already identified in these environments [24, 26, 27, 76, 102, 108].
floods, these floodplains tend to be elevated and uniform in topography and are covered pri-
marily with alluvial forest, oxbow and scroll bar lakes [29, 100]. Due to the high gradient and
sediment load of the rivers, they are subject to rapid lateral erosion and channel change [20]
which results in unstable levees penetrated by numerous scroll bar channels [29, 100]. These
channels allow exchange of water between river and floodplain, resulting in a high level of con-
nectivity throughout much of the hydrological cycle [29]. As with Balbina and Tucurui, then,
we expect a close correlation between changes in river stage variability and flood periods on
these floodplains. However, since higher elevations tend to predominate on the Andean flood-
plains, a reduction in mean stage height at high water could result in a much larger portion of
the floodplain being permanently isolated than was observed below Balbina and Tururui
dams. This could have profound impacts on the biota occupying these areas, especially on the
phenology of floodplain plants [37, 38] and the survival of animals that depend on these plants
[15, 32, 33]. The isolation of large parts of the floodplain may also promote human occupation
in these areas and accelerate rates of deforestation in alluvial forests.
yields estimated for the Andean reservoirs (Table 5) could compensate, in part, for the loss in
fish yields expected below the dams (Fig 6). The estimated fish yield in Manseriche reservoir
(15,200 tons km-2 y-1), for example, is similar to the average fish yield of the Loreto Region
(19,400 tons km-2 y-1). However, the impact on the livelihoods, economy and protein supply of
the populations downstream from the dams will likely be severe, and mercury contamination
both above and below the dams could create additional issues (see below).
Reservoir sedimentation
Besides the impacts on downstream sediment and nutrient supplies, reservoir sedimentation
can also affect dam operation, the quality of benthic habitats and the pattern of human occupa-
tion and activities in the reservoirs. Dam operation will be affected if a reservoir fills to the
point that sediment begins to enter the turbines. While this can be mitigated or delayed by
dredging and flow management, when sediment levels reach the turbine inflow it often marks
the end of the useful life of the power plant. If and when this occurs depends on the rate of fill-
ing (Table 6), the spatial distribution of sedimentation and the depth of the turbine inflow. For
the Manseriche and Inambari reservoirs where complete fill times were estimated at 6240 and
1225 years, respectively, sediment damage is unlikely to occur before other factors terminate
dam operations [112]. In contrast, the complete fill times for Angosta del Bala and Rositas res-
ervoirs, 106 and 126 years, respectively, are near the expected useful lives of most dams (50–
100 years [112]), and fill times to the turbine inflow are expected to be considerably shorter,
which could affect the technical viability of these dams.
Several environmental impacts linked to sedimentation must also be considered. Deposits
of fine riverine particulates will cover the coarse gravel substrates originally encountered on
the river bed, impacting the benthic fauna and fish which use this habitat for feeding and
spawning. Bottom sediments will accumulate first near the inflowing tributaries, where new
wetlands are expected to develop. These wetlands could provide new habitats for regional flora
and fauna, but could also provide new sources of greenhouse gases and sites for mercury meth-
ylation. Gold mining of alluvial deposits in the basin upstream from the dam site are currently
restricted to narrow floodplain environments, due to the high risk of mining in the fast flowing
river channels. The extensive deposits of alluvial sediments expected to accumulate near tribu-
tary mouths in the reservoir will provide an ideal environment for placer mining and is likely
to lead to an expansion of these activities in the region. The expansion of mining and other
activities upstream of the reservoir could increase the flux of sediments and associated mercury
to the reservoir, accelerating siltation and mercury contamination.
shown to contribute significantly to total emissions in other Amazonian dams [47, 86, 87, 88]
and should always be considered in environmental impact assessments of these systems.
Whether the values presented in Table 7 should be considered net or gross emissions will
depend on the expected changes in the regional carbon balance following impoundment. Net
emission, in this context, represents the net difference between the balance of CO2 equivalent
carbon in the reservoir landscape before and after impoundment. Most of the area to be inun-
dated by the four Andean reservoirs is currently occupied by tropical broadleaf forest, while a
smaller fraction is occupied by river channels and fluvial wetlands. A recent review of C mass
balance analyses for the lowland Amazon forest [114] concluded that this system is approxi-
mately carbon neutral. Several regional analyses have estimated high gross CO2 equivalent car-
bon emissions from river channels and wetlands in the lowland Amazon [115, 116, 117].
However when carbon uptake by aquatic plants was also considered in more complete mass
balance analyses [26, 118] these systems were also found to be approximately carbon neutral.
The errors involved in all of these mass balance analyses are too large to determine with confi-
dence whether these systems are carbon neutral, net sinks of net sources of carbon for the
atmosphere, and no similar mass balances are available for the forests and wetlands in the
Andean region. Based on the results for lowland regions cited above, it is likely that the forests
and wetlands currently occupying the areas of the projected Andean reservoirs are also close
to carbon neutral and that the emissions released from these areas once inundated (Table 7)
will represent net values. We do not consider in this analysis the assimilation of atmospheric
CO2 by the terrestrial forest during the period its biomass was growing, since this occurred
over millennia. CO2 equivalent carbon fluxes derived from the decomposition of this biomass
once submerged are therefore considered net emissions. We also ignore CO2 emissions associ-
ated with aquatic primary production in the reservoir, since these are balanced with recent
CO2 fixation. However methane emissions associated with aquatic production could contrib-
ute significantly to net CO2 equivalent emissions, due to the difference in global warming
potential of the fixed CO2 and released CH4 [85].
With greenhouse gas emissions rising globally and the consequences of global warming
becoming increasingly apparent, the choice of power generating technologies often involves a
comparison of Carbon Emission Factors (CEFs). Bosi [119] estimated CEFS for thermoelectric
power plants burning natural gas, fuel oil and coal in Brazil to be 0.115, 0.205 and 0.257 tons C
MWh -1, respectively. All of the hydroelectric dams considered here had CEFs below these fos-
sil fuel alternatives (Table 7), with the exception of the Manseriche, which had a predicted CEF
(0.206 tons C-CO2e MWh -1) similar to an oil-fired power plant and almost twice as high as a
gas-fired power plant. Considering the abundance of natural gas reserves in the Amazon
Region and the large environmental impacts associated with dams, alternative energy sources
should be considered in the case of Manseriche Dam.
Mercury dynamics
The historical pattern of mercury contamination in Cichla sp. in Balbina Reservoir (Fig 8) was
similar to that encountered for large predatory fish like Esox sp. in north temperate reservoirs,
with a gradual rise and fall of mercury during the first 10–25 years after impoundment [50, 51,
120]. Peak mercury levels in north temperate storage reservoirs were found to occur 3–13
years after impoundment with mercury concentrations returning to pre-impoundment levels
10–25 years after impoundment [50, 51, 120]. These trends have been attributed to a gradual
rise in the methylation and bioaccumulation of inorganic mercury, naturally present in these
systems, due to the slow decomposition of inundated terrestrial vegetation that generates
anoxic conditions favorable for methylation. As the terrestrial carbon stock is exhausted and
oxygen concentrations rise, the levels of methylation and contamination in aquatic biota
return to their pre-impoundment values. Mercury levels in Cichla sp. rose during the first ten
years after impoundment reaching a peak value of 0.65 μg g-1 between 1997 and 1998 (Fig 8).
The level then declined during the next 6 years and had still not returned to pre-impoundment
levels 25 years after impoundment [53]. The peak level of Hg observed in Cichla sp. exceeded
the value of 0.5 μg g-1 recommended by the WHO and the Brazilian federal government for
safe consumption.
Amazon reservoirs differ from north temperate impoundments in that they often have
commercial fisheries that exploit the large fish populations which commonly develop in these
systems. Fishing communities at Balbina and other Amazon reservoirs obtain most of their
protein from reservoir fish and are effectively the top predators in the reservoir food chain. It
is not surprising then that the historical trend in the levels of mercury in the hair of Balbina
fishermen’s wives was similar to those found in fish (Fig 8) with peak mercury levels in hair
occurring in 1991, just 1–2 years after the peak in fish values. Mercury concentrations in
hair were higher than those in fish reaching a maximum level of 7.5 μg g-1. Concentrations
exceeded the conservative reference limit of 1 μg g-1 established by the USEPA during the
entire study period and exceeded the reference limit of 6 μg g-1 established by the WHO
between 1997 and 2000, indicating an elevated health risk for this population.
Mercury contamination is also expected below hydroelectric reservoirs as anoxic hypolimi-
netic waters, rich in MeHg, are released through the turbines to the downstream river channel.
Kasper et al. [53] found elevated MeHg concentrations up to 200 km downstream from Bal-
bina Dam. Fish below Balbina and Samuel dams were shown to have higher levels of Hg con-
tamination than those found in the respective reservoirs [52, 53] due to the contaminating
effect of turbine discharge.
Similar patterns are expected in the six Andean reservoirs with some important differences.
All of these reservoirs will inundate extensive terrestrial forests which are expected to decom-
pose and create conditions conducive to mercury methylation for at least 10. During this
period, mercury levels are expected to increase in reservoir fish and in human populations that
consume these fish. However, unlike most reservoirs in the lowland Amazon and north tem-
perate regions, the anoxic conditions in the Andean reservoirs may continue well beyond the
point when terrestrial carbon sources are exhausted due to high levels of primary production
fueled by Andean nutrients. External nutrient loading is expected to be high and constant in
these systems, resulting in eutrophic reservoirs with permanently anoxic hypolimnia. These
stable anoxic environments will promote the methylation and bioaccumulation of mercury in
aquatic biota and fish-eating human populations upstream and downstream of the dams for
the useful life of the reservoir. Inputs of mercury from gold mining activities upstream of these
dam could also contribute to higher levels of contamination in these systems. The high sus-
tained fish yields predicted for these reservoirs (Table 5) are expected to attract fishermen
interested in exploiting this resource. These people are likely to consume large quantities of
reservoir fish and bioaccumulate significant levels of mercury. If the mercury levels in these
populations and in riverine communities downstream from the dam reach those encountered
in Balbina reservoir (Fig 8), they would represent a serious health risk, especially for children
and pregnant women.
Conclusions
The construction of six Andean dams will have extensive impacts on Amazon fluvial ecosys-
tems that need to be addressed in regional infrastructural development plans. Fig 9 summa-
rizes the principal impacts considered here, indicating their relative magnitude and
Fig 9. Summary: Expected environmental impacts above and below Andean dams.
https://doi.org/10.1371/journal.pone.0182254.g009
importance to ecosystem structure and functions. Understanding the full complexity of these
impacts and their economic costs should inform strategic discussion of whether alternative
energy sources could meet growing energy needs while preserving the critical natural
resources of the Amazon River basin.
Supporting information
S1 Fig. Relationship between particulate P and total suspended sediments (TSS) for the
Amazon mainstem and its principal Brazilian tributaries. Unpublished data from CAM-
REX Project, http://dx.doi.org/10.3334/ORNLDAAC/904.
(TIF)
S2 Fig. Relationship between % N in sediments and total suspended sediments (TSS) in the
Amazon mainstem and its principal Brazilian tributaries. Unpublished data from CAM-
REX Project, http://dx.doi.org/10.3334/ORNLDAAC/904.
(TIF)
S1 Table. Historical fisheries and flooding data for the Loreto fishing region of Peru.
Flooded areas simulated using the MGB-IPH Large Scale Numerical Hydrological Model,
developed specifically for the Amazon Basin [67, 68]. Fisheries data supplied by DIREPRO
(Dirección Regional de Producción, Peru). TY = total annual fish yield, tons y-1; YFZ = annual
fish yield for regularly exploited fishing zones, tons y-1; FZ = number of fishing zones. FZ was
assumed to represent fishing effort. Mifa = minimum annual flooded area, km2;
Mxfa = maximum annual flooded area, km2.
(XLSX)
Acknowledgments
The synthetic work for this paper was supported by the Science for Nature and People (SNAP)
sponsored by the National Center for Ecological Analysis and Synthesis (NCEAS), the Wildlife
Conservation Society (WCS) and the Nature Conservancy (TNC). At WCS we thank Cristián
Samper, John Robinson, Julie Kunen, Mariana Varese, Mariana Montoya, Carlos Durigan,
Guillermo Estupiñán, Micaela Varese, Natalia Piland and Sofia Baca; for SNAP workshop sup-
port we thank Charo Lanao; at TNC Craig Groves and Peter Kareiva; at NCEAS Frank Davis
and Lee Ann French. We thank D. Kasper, J.R.P. Peleja, E.M. Nakazono, A.P. Souza, J.B,
Rocha and D.R. Dietrich for help during the collection and analysis of fish and hair for mer-
cury at Balbina Reservoir and orientation. We also thank the Museu Paraense Emilio Goeldi
(MPEG) and the Instituto Nacional de Pesquisas da Amazônia (INPA) for their support. For
fisheries data we are indebted to the Dirección Regional de la Producción (DIREPRO), Peru.
SNAP funding was provided by the David and Lucile Packard Foundation (Grant # 2013–
38757 & #2014–39828), Ward Woods (Grant # 309519), the Wildlife Conservation Society
(WCS) and The Nature Conservancy (TNC). Field data reported for the first time was sup-
ported by the Gordon and Betty Moore Foundation (Grant 500) and the John D. and Cather-
ine T. MacArthur Foundation (Grant 84377). The Brazilian National Research Council
(CNPq) provided a productivity grant to B.R. Forsberg (Grant 309636/2011-6). J.M. Melack
received support from the US Department of Energy (Contract No. DE-0010620), NASA and
a Fulbright fellowship. From the above institutions, past or present, we thank Avecita Chic-
chón, Adrian Forsyth, Rosa Lemos da Sá and Enrique Ortiz.
Author Contributions
Conceptualization: Bruce R. Forsberg.
Data curation: Bruce R. Forsberg, Ronaldo B. Barthem, Michael Goulding, Rodrigo C. D.
Paiva, Mino V. Sorribas, Urbano L. Silva, Jr., Sabine Weisser.
Formal analysis: Bruce R. Forsberg, Thomas Dunne, Ronaldo B. Barthem, Rodrigo C. D.
Paiva, Mino V. Sorribas, Urbano L. Silva, Jr., Sabine Weisser.
Funding acquisition: Michael Goulding.
Investigation: Bruce R. Forsberg, John M. Melack.
Methodology: Bruce R. Forsberg, John M. Melack, Ronaldo B. Barthem, Rodrigo C. D. Paiva,
Mino V. Sorribas, Urbano L. Silva, Jr., Sabine Weisser.
Project administration: Michael Goulding.
Software: Rodrigo C. D. Paiva, Mino V. Sorribas.
Validation: Rodrigo C. D. Paiva, Mino V. Sorribas.
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